aunitaxys
Revue de Faunistique, Taxonomie et Systématique
morphologique et moléculaire
Volume 7 ISSN: 2269 - 6016
Numéro 4 Mars 2019 Dépét légal: Mars 2019
Faunitaxys
Revue de Faunistique, Taxonomie et Systématique
morphologique et moléculaire
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Imprimé le 20 mars 2019
Faunitaxys, 7(4), 2019: 1-25.
Review of the Phyllium Mliger, 1798 of Wallacea, with description of a
new subspecies from Morotai Island (Phasmatodea: Phylliidae: Phylliinae)
ROYCE T. CUMMING (1), STEPHANE LE TIRANT (2) & FRANK H. HENNEMANN (3)
(1) Entomology Department, San Diego Natural History Museum, POB 121390, Balboa Park, San Diego, California, United States,
92112-1390. Associate researcher for the Montréal Insectarium, Québec, Canada; H1X 2B2 - phylliidae.walkingleaf@gmail.com
- ZooBank : http://zoobank.org/6CA8501F-10BA-4E07-9BF4-65CFCE4E9E92
(2) Collection manager, Montréal Insectarium, 4581 rue Sherbrooke, Montréal, Québec, Canada, H1X 2B2 - sletirant@ville.montreal.qc.ca
- ZooBank : http://zoobank.org/A9391F8A-15D7-4D3B-9E3F-7123BA27EA2E
(3) Reiboldstrasse 11, 67251 Freinsheim, Germany- hennemann@phasmatodea.com
- ZooBank : http://zoobank.org/65 1 FCCFA-271B-48A3-A58E-A30FDC739493
Abstract. — The Phylliidae of Wallacea are here reviewed, with notes on species
distributions. Phyllium (Phyllium) tobeloense GroBer, 2007 is also here recorded as
having a new subspecies from Morotai Island, Phyllium (Phyllium) tobeloense
bhaskarai Cumming, Le Tirant, and Hennemann ssp. nov. based largely upon the
distinct egg morphology. To conclude a key to all known Phyllium from Wallacea is
presented with most features illustrated for ease in identification.
Cumming R. T., Le Tirant S. & Hennemann F. H., 2019. — Review of the Phyllium Illiger, 1798
of Wallacea, with description of a new subspecies from Morotai Island (Phasmatodea: Phylliidae:
ZooBank : http://zoobank.org/CEC3E930-3ESD-4B2A-AE4D-51A198FD9E9E
Keywords:
Phasmatodea ; Morotai ;
Phasmida ; tobeloense ;
Phylliidae ; bhaskarai ;
Phylliini ; description ;
Phyllium ; new subspecies ;
Wallacea ; distribution ; Phylliinae). Faunitaxys, 7(4) : 1-25.
Indonesia ; biogeography.
Introduction
Members of the genus Phyllium Illiger, 1798 are well known as
“Leaf Insects” or “Walking Leaves” and their still poorly
understood systematics, phylogeny, and distributions have been
studied more comprehensively during the past ten years. It has
been shown that the actual biodiversity of these fascinating
insects is far larger than previously suggested and several
papers by the authors and various colleagues have described 30
new species of Phyllium from throughout Southeast Asia since
2009 (Brock et al., 2018). The Phylliidae of Wallacea are
particularly interesting but have not yet been studied in any
detail, with only some of the species described before 2009
covered by Hennemann et al. (2009). In addition, two new
species from Wallacea have recently been described from the
islands of Peleng (Cumming & Teemsma, 2018) and Lombok
(Cumming, Valero & Teemsma, 2018). Examination of
extensive fresh material that has recently become available
from various islands of Wallacea has revealed interesting facts
concerning the synonymies and distributions of the species
concerned and are summarized in the present paper. This
includes a new subspecies from the Island of Morotai north of
Halmahera, Maluku Islands, which is described herein. Still
however, many questions concerning the true distributional
ranges of some species throughout the very complex Wallacea
subregion remain unanswered and the authors are aware of
further still undescribed species, that will hopefully sometime
become available for scientific recognition in order to clarify
the biogeographic and phylogenetic facts and help to complete
our knowledge of Wallacean Phylliidae.
Biogeography
Wallacea is a group of several thousand islands of various
sizes, which cover a total land area of roughly 338,500 km?
located between Sundaland in the west, the Philippines in the
north as well as New Guinea and Australia in the east and
southeast. Recent studies suggest that also the Philippines need
to be considered as part of the Wallacea (e.g. Vallejo, 2011), but
for covering the Phylliidae of this particular region Wallacea is
here treated in its traditional definition that excludes the
Philippines. All islands are located almost entirely within the
borders of Indonesia and include the island of Sulawesi, the
Maluku Islands (also called Moluccas), the Kai Islands, Banda
Islands and the Lesser Sunda Islands (except Bali). The largest
of these islands is Sulawesi. The Lesser Sunda Islands
attributed to the Wallacea are located south of Sulawesi and
include Lombok, Sumba, Sumbawa, Flores and Timor, as well
as several hundred smaller intervening islands. The Maluku
Islands include several hundred islands in the northern portion
of Wallacea, the largest being Halmahera in the north and
Seram in the south. The most northern islands of the Maluku
Islands are the Sangihe Islands and Talaud Islands, the most
southeastern ones are the Kai Islands and Banda Islands,
situated west of New Guinea.
Principally, Wallacea is a transition zone between the Oriental
region and the Australian region with the fauna showing
influences from both these regions (Dickerson, 1928). The
western border of Wallacea is represented by Wallace’s line
(named by Huxley in 1868), which runs between Borneo and
Sulawesi in the north and Lombok and Bali in the south (Fig.
1). The eastern border of Wallacea has been disputed by
2 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
geologists and biogeographists and attempts to draw a definite
line have produced a number of different lines (Simpson,
1977). The most accepted of these lines is Lyddeker’s line
which follows the Sahul shelf, that runs between Seram and the
Banda Islands in the west and New Guinea in the east (Fig. 1).
Another well-known and much discussed line is Weber’s line,
which roughly runs between Sulawesi and the Maluku Islands
(Gressitt, 1982). Mayr (1944) regarded Weber’s line as a line of
faunal balance, east of which the Papuan faunistic elements
have a numerical superiority over Oriental elements.
Wallacea is regarded as a biodiversity hotspot although the
subregion contains fewer known species overall compared to
e.g. Sundaland, the Philippines, or the Australian region.
However, Wallacea contains a very high percentage of endemic
species, i.e. species that are only found in Wallacea or on single
islands within this subregion, which is explained by several
factors. First of all, the subregion contains a large number of
different islands and geographic isolation is one of the major
factors which leads to diversification and formation of endemic
species. Wallace’s line marks the position of deep ocean straits
and separates shallow waters in the west from deep waters in
the east. The sea-level dropped by an estimated 120 meters
during the last ice-age and while the shallow waters in the west
disappeared and formed land connections between the Large
Sunda Islands (Sumatra and Java), Borneo, and the remaining
portions of Sundaland, there were no such land bridge
connections towards Wallacea.
Materials and Methods
Photos of specimens were taken by René Limoges of the
Montreal Insectarium (IMQC) using a Nikon D810 DSLR
camera with Nikon Micro-Nikkor 200mm f/4 lens on
Manfrotto 454 micrometric positioning sliding plate. Lighting
was provided by two Nikon SB-25 flash units with a Cameron
Digital diffusion photo box. Adobe Photoshop Elements 13 was
used as post processing software. Measurements of specimens
were made to the nearest 0.1 mm using digital calipers. The
Phyllium (Phyllium) tobeloense bhaskarai ssp. nov. holotype
and allotype specimens are deposited in the Montreal
Insectarium Quebec, Canada (IMQC) type collection.
Photographs of specimens within the Frank Hennemann
personal collection were taken by himself using a Nikon D7000
camera equipped with a Nikon DX AF-S Micro 40mm lens and
a wireless Nikon SU-800 dual speed light system. Background
lightning was provided by a 18W 6000K LED panel light plate.
The photographs of the female holotype Phyllium
(Pulchriphyllium) suzukii GroBer, 2008 was taken by Mandy
Schroter under direction of Stephan Blanke at the Senckenberg
German Entomological Institute Mtincheberg using a Nikon
D7200 digital camera and a Nikon Micro Nikkor 105mm f/2.8
G ED objective. Lightning was from the Yongnuo Digital
Speedlight YN 560 IV reflected by the inner surface of a
styrofoam box set up around the specimen. A grey card was
used for white balance. Composite images with an extended
depth of field were created using the software StackShot Macro
Rail Package (Cognisys Inc., U.S.A.) and Zerene Stacker
(release November 7, 2017; Zerene Systems LLC, U.S.A.).
Abbreviations
— AMSA: Australian Museum, Sydney, NSW / Australia.
—BPBM: Bernice P. Bishop Museum, Honolulu, Hawaii, U.S.A.
— BYU: Monte L. Bean Life Science Museum, Brigham Young
University, Provo, Utah / U.S.A.
—CASC: Califomia Academy of Sciences, San Francisco, California / U.S.A.
—IMQC: Insectarium de Montréal, Montréal, Québec / Canada.
—IRSN: Institut Royal des Sciences Naturelles, Brussels / Belgium.
—LEMQ: Lyman Entomological Museum, McGill University, Québec, Canada.
—MNHN: Museum National d'Histoire Naturelle, Paris / France.
— MNHU: Museum fiir Naturkunde der Humboldt-Universitét, Berlin /
Germany.
—MZSF: Museum Zoologique, Université de Strasbourg, Strasbourg/ France.
—NHMUK: The Natural History Museum, London / United Kingdom.
—NHMW: Naturhistorisches Museum Wien, Vienna / Austria.
—NMR: Natuurhistorisch Museum, Rotterdam / Netherlands.
—RMNE: Nationaal Natuurhistorisch Museum Naturalis, Leiden / Netherlands.
— SDEI: Senckenberg Deutsches Entomologisches Institut, Miincheberg /
Germany.
—SDNHM: San Diego Natural History Museum, San Diego, California / U.S.A.
—SMEM: Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt a. M. /
Germany.
—SMNS: Staatliches Museum ftir Naturkunde, Stuttgart / Germany.
—UCRC: University of California, Dept. of Entomology, Riverside, California /
USA.
— UUZM: Uppsala University Zoological Museum, Linnean collection,
Uppsala / Sweden.
—ZSMC: Zoologische Staatssammlung, Munich / Germany.
— AB: Private collection of Alexander Banko / Canada.
— EB: Private collection of Edy Bhaskara / Indonesia.
— FH: Private collection of Frank H. Hennemann / Germany.
—MO: Private collection of Maxime Ortiz / France.
— OC: Private collection of Oskar V. Conle / Germany.
—RTC: Private collection of Royce T. Cumming / U.S.A.
— SLT: Private collection of Stéphane Le Tirant / Canada.
—ST: Private collection of Sierra Teemsma / U.S.A.
—TM: Private collection of Tetsuo Miyashita / Japan.
Note on Locality Spelling Changes
With a number of historic data labels listed within, it is
important to note the various spellings of localities which have
changed throughout the years. Although not all those listed
below are used within this work, we have noted additional
spellings from throughout Wallacea to assist others who may be
reviewing historic Wallacean specimens.
Present Common Name = Known Historic Names
—Ambon Island = Ambonia / Amboina
—Bacan Island = Bachans / Bachians / Batchians
—Banggai Islands = Bangai / Bankei / Bangkei
—Buru Island = Bouru / Boro / Boeroe
—Buton Island = Button / Boeton / Butung
—Halmahera = Jilolo / Gilolo / Jailolo
~Kai / Kei Islands (both commonly used at present) = Key Islands
— Maluku Province = Moluccas / Muluku Province
—Sanana Island = Sula Besi / Xulla Besi
—Seram Island = Ceram / Seran / Serang
— Sulawesi = Celebes
—Taliabu Island Regency = Taliaboe
Treatment of Species
Subgenus Phyllium (Pulchriphyllium) Griffini, 1898
Type species: Phyllium pulchrifolium Audinet-Serville, 1838, 292, by
original designation.
Remarks. — Only two species currently in this subgenus and
attributed to the frondosum species group defined by
Faunitaxys, 7(4), 2019: 1-25.
-Wallace Line
“
_ Weber Line
; Lydekker Line
BORNEO
NEW
GUINEA
Fig. 1. Area defined as Wallacea (in green), bordered to the northwest by Wallace’s line and to the southeast by Lydekker’s line.
Morotai
Tobelo. Galela
{rondano Mtlbu
QPalolo
Sanana
Cungadial
5 P tly
Mamasa
eran
@imbon
P re y
BP. kovicum SP. sp. “Suton” BP. letirand BP. mamasnense >, tobaloanse bhaskare
Ys Krayicunn “ay SP ¥ eiranti ¥ in ¥ tobdlosnse bhaskaral
4
Fig. 2. Distribution map of the Phyllium species currently known within the boundaries of Wallacea. (Google Earth: Image
Landsat/ Copernicus: Data SIO, NOAA, U.S. Navy, NGA, GEBCO: Accessed December 13th, 2018.
4 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
Hennemann et al. (2009) are known from Wallacea at present.
One is only known from a unique specimen collected on the
island of Bacan west of Halmahera in the northern Maluku
Islands and the other species is not uncommon on the Kai
Islands in the very southeastern portion of Wallacea.
Additionally, both species are the only ones of the frondosum
species group that are found outside of mainland New Guinea,
and currently only known from female specimens.
Phyllium (Pulchriphyllium) suzukii GroBer, 2008
(Fig. 5, 7,9 & 61)
Phyllium (Pulchriphyllium) suzukii GréBer, 2008: 137, fig. 171-172.
Holotype, 2: Phyllium (Pu.) suzukii det. GréBer, Molukken, Ins. Bacan,
9.2006; Phyllium (Pu.) suzukii GréBer sp. nov. Holotypus, det. D. GréBer
9.2006; DEI Hemimetabola #100121 [SDEI].
Distribution.
— INDONESIA: North Maluku Province, South Halmahera
Regency, Bacan Island [SDEI].
Discussion. — Phyllium suzukii is one of only two species from
the frondosum species group found outside of mainland New
Guinea (the other being Phyllium keyicum Karney, 1914
discussed below) and is currently the only known species of
Phyllium recorded from Bacan Island. This species is currently
only known from the single holotype female held within the
SDEI. This species is most characterized by the unique
mesopraescutum which is 3.7 times wider than long (Fig. 9, the
most extreme width to length ratio of known Phyllium females)
as well as an abdomen with prominent, posteriorly directed
projections on segments VI-VIII giving the abdomen a unique
overall shape (Fig. 5); the outer margins of segments VII and
VII posteriorly project beyond the apex of the anal segment.
With a body length of only 57.6 mm it is the smallest of all
know representatives of the Phylliidae within the boundaries of
Wallacea. The egg was illustrated by GréBer (2008: 137, fig.
171) and is easily recognized by the unusually ovoid overall
shape and large roughly circular impressions of the capsule.
Phyllium (Pulchriphyllium) keyicum Karny, 1914
(Fig. 3, 4, 6 & 8)
Phyllium keyicum Karny, 1914: 7, fig. 5. Holotype, 9: Kei Islands,
Toeal Island [depository unknown].
= Phyllium insulanicum Werner, 1922: 126. Holotype, 2: Toeal, Kei-
eilanden, J.H. Jurriaanse 1920 [RMNH]. (Synonymised by
Hennemann et al., 2009: 65)
= Phyllium indicum Ginther, 1929: 614 (Misspelling of insulanicum).
Distribution.
— INDONESIA: Maluku Province, Southeast Maluku Regency,
Kai Islands [NHMUK, NHMW, NMR, RMNH, SMNS, FH].
Discussion. — A detailed description of the females of this
species and the synonymous Ph. insulanicum was presented by
Willemse (1947). Since the holotype female of Ph. keyicum is
not traced and presumed lost (Otte & Brock, 2005: 274), it may
become necessary to designate a neotype. The egg was
illustrated by Gr6Ber (2008: 123, fig. 146).
Despite the Kai islands being rather remote, there have been a
number of female specimens collected over the years.
Unfortunately despite the female being not uncommon, the
male Ph. keyicum is still unknown. This species is most notable
for its profemoral exterior lobe with a posterior margin that is
straight or very nearly so (Fig. 6, black line), this is the only
species within the frondosum species group with this feature.
Ph. keyicum also has the unique morphology of a single
posteromedial tubercle on the head capsule versus other
members of the frondosum species group which have the
tubercle split into two. Their most variable morphological
feature is the terminal abdominal segments which can range
from nearly straight across (Fig. 4) to more pronounced lobes
(Fig. 3) and the full range between these two extremes. This
intraspecific variability led to the description of Phyllium
insulanicum Werner, 1922, based on the more strongly lobed
females. Willemse (1947) extensively described and compared
a series of specimens from the Kai Islands and from his
thorough review it appears as though the variation in the
abdominal lobes is the only notable feature between the two
taxa. It was not until Hennemann et al. (2009: 65) correctly
synominized the Ph. insulanicum with Ph. keyicum, leaving
there only a single species of Phyllium known from the Kai
Islands. Variation in abdominal shape is now well known as a
common occurrence in many species of Phyllium and not a
feature that should singularly be relied upon for differentiation.
With a body length of 70-79 mm females of this species are
considerably larger than the related Ph. suzukii from Bacan
Island.
Subgenus Phyllium (Phyllium) Iliger, 1798
Type species: Gryllis (Mantis) siccifolius Linnaeus, 1758: 425, by
monotypy.
Remarks. — The seven species known to occur throughout
Wallacea belong to two species groups defined by Hennemann
et al. (2009). While only the Sulwesian Ph. celebicum de Haan,
1842 belongs to the celebicum species group all other known
species fall within the siccifolium species group.
Phyllium (Phyllium) celebicum de Haan, 1842
(Fig. 10-16, 81 & 82)
Phasma (Phyllium) celebicum de Haan, 1842: 111. Holotype, 9:
Celebes, Tondano [RMNH - not traced].
Distribution.
— INDONESIA: North Sulawesi Province, Minahasa Regency,
Tondano [RMNH - type locality].
— Central Sulawesi Province, Sigi Regency, Palolo [RC].
— South Sulawesi Province, North Luwu Regency, Bungadidi
[FH, OC, SLT].
— South Sulawesi Province, North Luwu Regency, Tiulapolu [FH, RC].
— Southeast Sulawesi Province, Buton Regency, Buton Island [OC].
Discussion. — The female holotype was first reported lost in
Hennemann et al. (2009: 15) after their extensive search within
the RMNH collection in April 2006. Examination of the
presumed holotype in the same collection (Bragg, 1996: 110)
has revealed a female clearly not celebicum as originally
described by de Haan (1842: 111) or illustrated by Willemse
(1947: 319, fig. 3). The concerned specimen has features much
more agreeing with Ph. tobeloense based on the thorax and
profemoral lobes and does not represent the lost Ph. celebicum
holotype.
True Ph. celebicum is discussed in depth in Hennemann et al.
(2009: 11) where the authors provided illustrations, a detailed
redescription of the female, descriptions of the previously
unknown males and eggs, summarized the intraspecific
variability, and discussed the numerous misidentifications over
Faunitaxys, 7(4), 2019: 1 — 25.
10 mm
10mm
Fig. 3-9. Females, frondosum species group, dorsal view. - 3: Phyllium keyicum [FH 0083-1]. - 4: Phyllium keyicum [NHMUK 012496951 (Kei Is.,
1923)]. - 5: Phyllium suzukii GréBer, 2008 holotype [SDEI]. - 6: Left profemor Phyllium keyicum; line illustrating the straight profemoral exterior
lobe proximal edge, a unique feature to this species [NHMUK 012496951 (Kei Is., 1923)]. - 7: Phyllium suzukii holotype; line illustrating the curved
profemoral exterior lobe proximal edge, and the arrow pointing out the protibial exterior spur, both features clearly contrasting Ph. suzukii from
Ph. keyicum [SDEI]. - 8: Mesopraescutum and mesopleurae Phyllium keyicum [NHMUK 012496951 (Kei Is., 1923)]. - 9: Mesopraescutum and
mesopleurae Phyllium suzukii holotype [SDEI].
the years. Phyllium celebicum is currently the only known
member of the celebicum species group from Wallacea. This
species group is characterized by males with profemoral lobes
which are broader than the interior lobes (Fig. 16), and females
which have developed alae (length 26.0-32.0 mm, fig. 11), both
features which easily separate them from any other Phyllium of
Wallacea.
Phyllium celebicum is currently only known from throughout
Sulawesi and from nearby Buton Island. The single female
specimen from the island of Ambon in the MNHU listed in
Hennemann, ef al., (2009: 11) has been reexamined and found
to not represent Ph. celebicum. Although very similar, this
female specimen has several features which differentiate it
from Ph. celebicum such as: a subgenital plate that is long and
slender almost reaching the apex of the abdomen (in PA.
celebicum the subgenital plate it short and stout, only just
reaching to terminal segment X); mesopleurae which reach
nearly to the anterior of the mesopraescutum (in Ph. celebicum
the mesopleurae are distinctly shorter reaching only half way
through the mesopraescutum); and alae which are notably
longer than any examined Ph. celebicum females (49.0 mm in
the MNHU specimen versus 26.0-32.0 mm in Ph. celebicum,
see Hennemann, ef al., 2009: 16). The vague historic collection
data of “Amboina” unfortunately cannot be trusted as truly
coming from the island of Ambon and must be taken with little
confidence. Realistically this celebicum species group member
likely represents one of the many poorly described species from
mainland Asia.
Body lengths: 9 9 75.5-92.0 mm, 34 62.0-66.0 mm.
6 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
Fig. 10-16. Phyllium celebicum dorsal. - 10-11: Females; note the well developed alae which help to characterize the celebicum species group
[FH 0634-5 (left) and 0634-1 (right)]. - 12: Male [FH 0634-4]. - 13-15: Female left profemor showing intraspecies variability [FH 0634-6, 0634-2,
0634-1]. - 16: Male left profemor; note the exterior lobe with its strong angle and notably larger than the interior lobe, a feature which defines
the celebicum species group [FH 0634-4].
Phyllium (Phyllium) conlei Curmming, Valero, & Teemsma, 2018
(Fig. 23 & 44)
Phyllium (Phyllium) conlei Cumming, Valero, & Teemsma, 2018: 3, fig.
3a-i. Holotype, 4: Indonesien, Lombok, XII.2012 [ZSMC].
Distribution.
— INDONESIA: West Nusa Tenggara Province, Lombok Island
[ZSMC].
Discussion. — Currently only known from the unique holotype
male deposited in the State Zoological Collection of Munich
(ZSMC) originally from the collection of Oskar Conle
(Germany). Cumming, Valero, & Teemsma (2018) discuss the
lack of knowledge for the Phylliidae of the Lesser Sunda Islands
and described Ph. conlei as the first unique species to the island
chain. With the unique male at only 47.5mm in length, Ph.
conlei is one of the smaller species of Phyllium. Nearby species
also with males of a similar size range are Phyllium jacobsoni
Rehn & Rehn, 1934: 42.5-50.4 mm, Java (Hennemann et al.,
2009), Phyllium hausleithneri Brock, 1999: 51.4-57.8 mm,
West Malaysia and Sumatra (Cumming, Le Tirant, & Teemsma,
2018), and Phyllium chengiae Seow-Choen, 2017 (53 mm,
Borneo). From all species of a similar size, Ph. conlei can be
differentiated by the slender abdomen with a width only about
30% of the total abdominal length versus the others with broad
spade shaped abdomens 40-55%.
Faunitaxys, 7(4), 2019: 1 — 25.
Fig. 17-22. Females, dorsal. - 17: Phyllium mamasaense holotype [SDEI]. 18-19: Phyllium letiranti green and brown colorforms [SLT coll.]. - 20:
Phyllium siccifolium holotype [UUZM]. - 21: Phyllium siccifolium from Seram Island [FH 1042-1]. - 22: Phyllium tobeloense tobeloense from
Halmahera Island, Mount Ibu [FH 0657-7].
Phyllium (Phyllium) sp. “Buton”
(Fig. 24 & 45)
Distribution.
— INDONESIA: Southeast Sulawesi Province, Buton Regency,
Buton Island [FH].
Discussion. — Known only from a single male within the
Hennemann private collection (FH 0673-1) this specimen was
the most difficult to place, with a near perfect match within the
intraspecies morphological range of Ph. hausleithneri Brock,
1999 and Ph. jacobsoni Rehn and Rehn, 1934, both species
found outside of Wallacea. Finding either species on Buton (far
from their known distributions outside of Wallacea) would be
very surprising and would challenge the well-defined
biogeographic boundary lines of Wallacea. The authors are
confident in the collection data, but, mistakes by collectors or
specimen dealers are a possibility and we cannot ignore the fact
that this could simply be a clerical error. Additionally, the
8 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
Fig. 23-29. Males, dorsal. - 23: Phyllium conlei holotype [ZSMC]. - 24: Phyllium sp. “Buton” [FH 0673-1]. - 25: Phyllium tobeloense tobeloense from
Halmahera Island, Mount Ibu [FH 0657-9], note the antennae not reaching the apex of the resting tegmina. - 26: Phyllium tobeloense tobeloense captive
reared from Halmahera Island, Galela [FH 0657-13]. - 27: Phyllium letiranti from Peleng Island [SLT coll.]. - 28: Phyllium letiranti from Sanana Island,
Waifara [FH 1008-10] note the antennae exceeding the apex of the resting tegmina. - 29: Phyllium siccifolium from Ambon Island [NHMUK].
Faunitaxys, 7(4), 2019: 1 — 25.
Fig. 30-45. Profemora, dorsal. - 30: Female Phyllium siccifolium holotype [UUZM]. - 31: Female Phyllium siccifolium [FH 1042-1]. - 32: Male Phyllium
siccifolium [NHMUK]. - 33: Female Phyllium mamasaense holotype [SDEI]. - 34-36: Female Phyllium letiranti showing intraspecies variation in color
and lobe serration. - 37: Male Phyllium letiranti [SLT coll.]. - 38-41: Females Phyllium tobeloense bhaskarai ssp. nov. paratypes, showing remarkable
intraspecies variation in color and lobe serration [FH 1077-3, 1077-1, 1077-2. 1077-6]. - 42: Male Phyllium tobeloense bhaskarai ssp. nov. allotype
[IMQC]. - 43: Male Phyllium tobeloense bhaskarai ssp. nov. paratype [FH 1077-21]. - 44: Male Phyllium conlei holotype [ZSMC]. - 45: Male Phyllium
sp. “Buton” [FH 0673-1].
female and egg from this “Buton” species are not yet known,
leaving the possibility that clear morphological features
identifying it as a new taxon might simply only be visible in the
female or egg morphology. With Ph. tobeloense bhaskarai ssp.
nov. only easily separated from the nominate subspecies by the
unique egg morphology, it is very possible that this male could
represent a new species or subspecies, only recognizable by
female or egg morphology. Therefore, with a lack of significant
morphological features to base a new taxon on, we simply refer
to this species as Phyllium (Phyllium) sp. “Buton” throughout
this work. Body length 48.5 mm.
Phyllium (Phyllium) letiranti Cumming & Teemsma, 2018
(Fig. 18, 19, 27, 28, 34-37, 50, 51, 54, 87, 88, 91 & 96-98)
Phyllium (Phyllium) letiranti Cumming & Teemsma, 2018: 2, fig. 1A-
D, 2B-D, F-G, 3A-E, 4A-D, 5A-D, 6A-C. Holotype, 9: Indonesia,
Peleng Island, Tinagkung Utara, near Luksagu Village: December,
2016: ~60 m. elv. 1°17’ S 123°25.4' E [IMQC]; Paratypes [IMQC,
SDNHM, CASC, SDEI, RC, ST, SLT, and FH].
Phyllium (Phyllium) siccifolium, Klante, 1976: 65, 76, fig. 10, (in
part).
Phyllium (Phyllium) siccifolium, Hennemann et al., 2009: 58 (in part).
10 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
Fig. 46-50. Females, ventral view of coxae. - 46: Phyllium mamasaense holotype, showing black interior mark on meso- and metacoxae
[SDEI]. - 47: Phyllium tobeloense tobeloense, showing metacoxae only with a black interior marking, mesocoxae lacking colored mark [FH 0657-3].
- 48: Phyllium tobeloense bhaskarai ssp. nov. paratype, showing metacoxae only with a black interior marking, mesocoxae lacking colored mark
[FH 1077-2]. - 49: Phyllium siccifolium, mesocoxae with black interior marking [NHMUK]. - 50: Phyllium letiranti live female bred by Maxime
Ortiz (France) showing lack of black coxae markings and the vibrant orange coxae color (© Maxime Ortiz).
Distribution.
— INDONESIA: Central Sulawesi Province, Banggai Regency,
Peleng Island [RC, FH, ST, SLT, CASC, IMQC, SDNHM];
Banggai Islands [MNHU].
— INDONESIA: North Maluku Province, Sula Islands Regency,
Sanana Island [FH - distribution expansion].
Discussion. — Only recently collected (2016) and described
(2018), this species has become widely available for sale as
dried specimens and has entered the phasmid breeding
community. In captivity it has been described as a relatively
easy species to raise with a trend to produce a wide variety of
female colorforms (Fig. 96-98). Originally described from
Peleng Island, the authors have received photos of Phyllium
from the nearby coast of Sulawesi (Central Sulawesi Province,
East Peninsula) which appear very similar in morphology to
Ph. letiranti, but without high resolution photos or specimens
to review, a confident identification is not possible at this time.
It would not be surprising to find Ph. letiranti on the Sulawesi
coast near Peleng Island but with the similar species Ph.
mamasaense also found on Sulawesi, from photos alone an
identification cannot be confirmed. Hopefully future collecting
on Sulawesi can bring to light the distribution of Ph. letiranti
with more clarity.
A range expansion can however be made for Sanana Island
(Sula Islands) as represented by a male specimen from the
Hennemann private collection (Fig. 28, FH 1008-10). With the
island of Sanana known to have PA. letiranti, the female from
“Taliaboe” (Taliabu) in the RMNH illustrated in Klante (1976)
and the female from the MNHU from “Bangkei” (Banggai)
discussed in Hennemann, et al., (2009) likely also represents
Ph. letiranti. Examination of the specimen within the MNHU
does reveal a female which matches well with slender PA.
letiranti females from Peleng Island.
The two easiest diagnostic features to differentiate female Ph.
letiranti from Ph. siccifolium are the ventral color of the coxae
(Ph. siccifolium having distinct black markings and Ph.
letiranti without) and the number of antennal segments (nine in
Ph. siccifolium and ten in Ph. letiranti). Unfortunately, in
Klante (1976) neither feature was illustrated clearly and
Faunitaxys, 7(4), 2019: 1 — 25. 11
therefore a thorough examination of the specimen in question
must eventually be done to confirm this identification.
With the Sula Islands situated just to the east of the Banggai
Islands, this distribution expansion is not surprising and opens
the possibility that this species can also be found on the other
Banggai or other Sula Islands.
Body lengths: 9 9 86.6-91.2 mm, 33 57.6-62.2 mm.
Phyllium (Phyllium) mamasaense GroBer, 2008
(Fig. 17, 33, 46, 68, 89 & 90)
Phyllium (Phyllium) mamasaense Groer, 2008: 124, fig. 147-148.
Holotype, 2: Phyllium (Ph.). mamasaense det. GrofBer, Sulawesi,
Mamasa, 9.2006; Phyllium (Ph.) mamasaense GrdBer sp. nov.
Holotypus, det. D. Gréer 9.2006; DEI Hemimetabola #100116
[SDE]].
Distribution.
— INDONESIA: West Sulawesi Province, Mamasa Regency,
Mamasa District [SDEI].
Discussion. — Only known from very few female specimens
collected over the last ten years from the type locality of
Mamasa, Sulawesi (Fig. 2). This species has the unique
characteristic of having meso- and metacoxae with black
interior markings on the ventral surface (Fig. 46) a feature
within Wallacea currently only known from Ph. siccifolium
(Fig. 49), or only partially seen in Ph. tobeloense (Fig. 47, only
metacoxae with black marking). Interestingly, all females
observed (although not a wide sample size) have had a very
steady morphology with abdominal segments VII and VIII
consistently roundly angular (Fig. 68) and the profemoral
interior lobe with two large prominent teeth (as well as several
small minor teeth) (Fig. 33). These features are variable among
certain other Phyllium females and we look forward to
hopefully reviewing more material in the future as well as
describing the currently unknown male once it is identified.
The large eggs are very distinctive (Fig. 89 & 90) and differ
from those of all other Wallacean Phylliidae by the feather-like
fringes of the capsule forming several parallel longitudinal
ridges and the the very large cup-shaped operculum.
Body length 93.5-105.0 mm.
Phyllium (Phyllium) siccifolium (Linnaeus, 1758)
(Fig. 20, 21, 29, 30-32, 49, 52, 55, 63 & 69)
Gryllus (Mantis) siccifolium Linné, 1758: 425, No. 3. Holotype, 9:
Indes, Gustav IV Adolph; Gryllus (Mantis) siccifolius type; Coll. n°
1818 [UUZM].
= Phyllium brevicorne Latreille, 1806: 272. (Synonymised by Gray, 1835: 30).
= Phasma chlorophylla Stoll, 1813, 69, pl. 23: 89. (Synonymised by
Redtenbacher, 1906: 176).
= Phasma citrifolium Lichtenstein, 1796: 78. (Synonymised by Gray,
1835: 30).
= Phasma donovani Gray, 1835: 31. (Synonymised by Redtenbacher,
1906: 176).
= Phasma foliatus Perry, 1810: plate 24. (Synonymised by Westwood,
1859: 172).
= Phasma gorgon Gray, 1835: 31. (Synonymised by Westwood, 1859:
172).
= Phyllium stollii Le Peletier de Saint Fargeau & Serville, 1827: 115.
(Unnecessary replacement name for Phasma chlorophylla Stoll, 1813).
[Not: Phyllium (Phyllium) tobeloense GréBer, 2007: 15. Erroneous
synonym of Hennemann et al., 2009: 53].
Distribution.
— INDONESIA: Maluku Province, Ambon [MNHU, NHMUKkK,
RMNH, UUZM], Seram [NHMUK, FH] and Buru Islands [NHMW].
Discussion. — As the type species for the family, this species is
one of great importance but unfortunately it is also one of the
many poorly known species because of the lack of available
material in modern times. The holotype female (Fig. 20) was
redescribed in Hennemann ef al. (2009: 55) as well as a male
specimen from the island of Ambon from NHMUK (Fig. 29).
These authors also discussed the numerous synonyms and
provided information of the intraspecific variability. The exact
type-locality of Ph. siccifolium originally stated to be “Habitat
in Indiis” by Linnaeus (1758: 425) was discussed in detail in
Hennemann et al. (2009: 57). These authors stated the holotype
female in the UUZM collection to be most likely from the
island of Ambon.
In addition to the island of Ambon, various collections
examined over the years have revealed numerous females from
the islands of Seram and Buru which match well with the
morphology of the holotype. The likely misidentification by
Klante (1976) discussed above from Taliabu Island is probably
the morphologically similar Ph. letiranti known from the
Banggai and Sula Islands, leaving true Ph. siccifolium
restricted to east of Weber’s Line (Fig. 1) and only known from
Ambon, Seram, and Buru. Unfortunately, the male of this
species is little known and the eggs are still unknown for Ph.
siccifolium. With egg morphology so drastically different
between PA. letiranti and Ph. tobeloense hopefully future fresh
material will reveal the eggs of Ph. siccifolium and their
morphology should shed light on the relationship to the other
similar species.
Body lengths: 9 89.0-94.0 mm, 34 67.0-73.0 mm.
Phyllium tobeloense Phyllium siccifolium
Ventral coxae color
Profemoral exterior
lobe*
2 (Fig. 38-41)
Antennae Length
Metacoxae with distinct black interior marking,
Q mesocoxae slightly orange or light green, no
black interior mark (Fig. 47 & 48)
Thin and smoothly arcing, greatest width only
1.5 to 2 times the width of the profemora shaft
Short, not projecting beyond the apex of the
oes resting tegmina
(Fig. 25)
Mesocoxae and metacoxae and with distinct black
interior marking
(Fig. 49)
Broad and slightly angular, greatest width 2.5 to 3
times the width of the profemora shaft
(Fig. 30 & 31)
Long, reaching or projecting beyond the apex of the
resting tegmina
(Fig. 29)
Table 1. Summary of comparison between Phyllium tobeloense and Phyllium siccifolium females and males.
* Measured at the proximal edge of the profemoral interior lobe. ** When antennae are held at rest back along the body.
12 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
twos [2 [ws [nme [wen
[—tenmorane fp esp fam
Table 2. Measurements of Phyllium (Phyllium) tobeloense bhaskarai ssp. nov., holotype female and allotype male. For
the paratype material only the smallest and largest in the series were measured to give a range. All measurements made
to the nearest 0.1mm.
* Including cerci and head, excluding antennae.
** Only measured on paratype specimen RC 18-421 which measures 101.2 mm in length.
Phyllium (Phyllium) tobeloense tobeloense Groer, 2007
(Fig. 22, 25, 26, 47, 64, 66, 83, 84, 92, 99 & 100)
Phyllium (Phyllium) tobeloense Gréer, 2007: 15, fig. 1-4. Holotype,
®: Phyllium (Ph.) tobeloense det. GréBer, Molukken, Halmahera,
Tobelo, 8.1998 [SDE]T].
Distribution.
— INDONESIA: North Maluku Province, North Halmahera
Regency, Tobelo [SDEI, FH, OC - type locality], Mount Ibu [FH,
RC, SLT] and Galela [FH].
Discussion. — Originally collected in Tobelo, Halmahera, this
species has been widely bred by enthusiasts and like PA.
letiranti has yielded a number of beautiful female colorforms
(Fig. 99 & 100). This species was briefly disputed in
Hennemann et al. (2009) where it was presented as a junior
synonym for Ph. siccifolium until GréBer (2011) where it was
reestablished. Here we agree with the reestablishment of Ph.
tobeloense as a valid species and discuss the morphological
reasoning for our decision.
Although very similar in morphology, when a review of a
significant series is examined these two species can be
separated by the following characteristics. The first feature for
differentiating females is the ventral color of the coxae, with
Ph. tobeloense having a black marking only on the metacoxae,
not on the mesocoxae (Fig. 47 & 48) versus Ph. siccifolium
with both meso- and metacoxae with black markings. Also for
females, the width of the profemoral exterior lobe is
consistently different with Ph. tobeloense having a more
slender lobe, no more than two times the width of the
profemoral shaft (Fig. 38-41), as compared to Ph. siccifolium
which always has a profemoral exterior lobe that is notably
wider (2.5 to 3 times) than the width of the profemoral shaft
(Fig. 30 & 31). For males, the antennae immediately separate
the two species, with Ph. tobeloense with shorter antennae, not
longer than tegmina length when at rest (Fig. 25), versus Ph.
siccifolium males which have longer antennae which reach to
or past the apex of the folder tegmina. These distinguishing
features are presented in summary in table 1. These features
have proven to be consistent across all specimens examined of
both populations, including extreme variations within each
population.
Body lengths: 29 89.0-101.0 mm, 33’ 59.0-62.0 mm.
Phyllium (Phyllium) tobeloense bhaskarai Cumming, Le
Tirant, and Hennemann, ssp. nov.
(Fig. 38-41, 48, 53, 56-58, 65, 67, 70-77, 79, 80, 85, 86, 101-103)
ZooBank : http://zoobank.org/4ED84870-BA6D-4546-
B518-278B66EEAF4D
Holotype: 9: INDONESIA, Morotai island, North Maluku, Pulau
Morotai Regency, Daeo village, XI-XII.2018 [IMQC].
Allotype: 3: INDONESIA, Morotai island, North Maluku, Pulau
Morotai Regency, Daeo village, XI-XII.2018 [IMQC].
Paratypes: [103 33, 90 2°, 46 eggs], INDONESIA, Morotai island,
North Maluku, Pulau Morotai Regency, Daeo Majiko village, XI-XII.
2018.
Depositions.
-14,2 99, 1 egg [AMSA]
-1¢,1 9 [BPBM]
-13,2 29, 1 egg [BYU]
Faunitaxys, 7(4), 2019: 1 — 25. 13
-2 88,2 22, 1 egg [CASC]
~5 34, 18 29, 10 eggs [IMQC]
~163,3 29 [IRSN]
~138,19 [LEMQ]
~2 33,2 29, 2 eggs [MNHN]
~13,3 29 [MNHU]
~163,1 9 [MZSF]
~2 33,2 29, 2 eggs [NHMUK]
~2 34,3 299° [NHMW]
~163,3 29 [RMNH]
-2 8,2 29 [SDE]
~2 83,2 29, 1 egg [SDNHM]
-13,3 99 [SMFM]
-13,2 99, 1 egg [UCRC]
-~2 83,3 29, 1 (penultimate instar) [ZSMC]
~13,1 9 [coll. AB]
-1 8,2 2 [coll. EB]
—15 33, 20 29, 12 eggs [coll. FH, No’s 1077-1 to 35, E]
-~1 4,1 [coll. MO]
~5 34,5 29 [coll. OC]
~6 64,7 29, 5 eggs [coll. RTC 18-421, 18-422, & 19-087 to 19-102]
~20 33, 20 99, 5 eggs [coll. SLT]
~1 4,1 9, 1 egg [coll. ST]
~1,1 9 [coll. TM].
Distribution.
— INDONESIA: North Maluku Province, Morotai Island
Regency.
Remarks. — Currently only known from Morotai Island, Indonesia.
Morotai is separated from northem Halmahera by the Morotai Strait,
a stretch of water only about 10 kilometres (6.2 mi) wide. The only
other island near Morotai is Bau Island, a small island on the west
coast. It is possible that Phyllium (Phyllium) tobeloense bhaskarai
ssp. nov. may also be found on Bau Island, due to its close
proximity but currently no specimen records exist.
Discussion. — This new population has only recently been
discovered (November, 2018 by Alim Yasin) on Morotai and
the first impression was that this was likely a range expansion
for Phyllium tobeloense until photos of the eggs were received.
Immediately it became apparent that this Morotai population
was not identical to the Halmaheran population. The decision to
classify this Morotai population a subspecies of Phyllium
tobeloense and not full species status is tentative but was based
on a number of observations. Upcoming genetic analysis will
help reveal the extent of the variability between the
populations, but our present morphological review suggests a
very close relationship. First is the incredible similarity of the
adult morphology between the two populations. With the
Halmaheran population rather popular in both phasmid
breeding communities and commonly bred for commercial
export as dried specimens, a wide review of the intraspecies
variation has been possible. This has revealed that the females
of this species have moderate variability in the profemoral
interior lobe, and slight variability in abdominal segment VIII.
Captive bred specimens have also resulted in a significant
variety of color forms (Fig. 99 & 100) and a variety of major
and minor profemoral interior lobes, similar to those seen in
exported dried specimens. With this significant view into the
intraspecies variation of males, females, and eggs, the egg
morphology differences between the Morotai and Halmahera
populations can easily be identified as not simply intraspecies
variation and is discussed in detail below. With egg
morphology one of the only concrete features to allow
differentiation between the populations, and the obvious
similarity between the population’s eggs, we consider this
Morotai population an offshoot of the mainland Halmahera
population and classify it as a subspecies.
An additional reason for our decision to classify the Morotai
population as a subspecies is the significant number of other
insect species which follow this trend. Frequently, the Morotai
populations are subspecies with sibling subspecies from the
other northern Maluku Islands. For example, Wallace’s golden
birdwing butterfly Ornithoptera croesus Wallace, 1859 is
endemic to the northern Maluku islands, with five total
subspecies distributed throughout these islands. In particular,
from Halmahera there is Ornithoptera croesus lydius (Felder &
Felder, 1865) and from Morotai Ornithoptera croesus toeantei
Parrot & Schmid, 1984 . Interesting, these butterfly subspecies
parallel our own Phyllium tobeloense subspecies as the females
are more easily morphologically separated than males are, with
males of these two-island populations near impossible to
differentiate (pers. comm. Gilles Delisle, birdwing butterfly
expert). As discussed in more detail below, we were able to find
consistent morphological differences between the females and
eggs of these two populations of Phyllium tobeloense but
between males no reliable features could be found.
Differentiation. — Phyllium tobeloense bhaskarai ssp. nov.
may be separated from the nominate subspecies from
Halmahera by only a few subtle morphological features. This
Morotai population is the first subspecies described for
Phyllium tobeloense GréBer, 2007 and is only easily separated
from the nominate by the egg morphology.
The most easily seen morphological feature separating the eggs
of the two subspecies is the overall shape when viewed
laterally. The nominate subspecies has eggs that are
asymmetrical with a reniform (kidney shaped) (Fig. 83)
whereas the new subspecies has eggs which are ovoid in shape
and almost symmetrical (Fig. 85). Also, in the nominate
subspecies the long pinnae along the margins are not fully
formed around the entire capsule with the area opposite the
micropylar plate and below the operculum bald (Fig. 83). In the
new subspecies the long pinnae along the margins are fully
formed around the entire capsule, with no bald portion opposite
of the micropylar plate below the operculum (Fig. 85).
Adults of the two subspecies are much more difficult to
differentiate morphologically with only subtle differences
found when examining a large series of both populations. The
only consistent feature observed on the females were the length
of the subgenital plate. In the nominate subspecies the
subgenital plate was stouter, protruding only about half the
length of sternite X, leaving a significant amount of the
gonapophyses exposed (Fig. 64), and for the new subspecies
the subgenital plate was longer, about three-quarter the length
of sternite X, leaving only the tips of the gonapophyses
exposed (Fig. 65).
After an extensive search for a consistent feature to
differentiate males of these two populations, none was found
that was reliable enough to be useful in a morphological key.
With this species so common thus allowing the intraspecies
variation to be well known, our conclusion is that it is likely
that only molecular comparison can confidently differentiate
these two populations males.
Habitat and Host Plant. — The local collectors were kind
enough to share a number of photos of live specimens in the
wild on their host plant (Fig. 78-80). The photos of this host
plant were shared with Binjie Ge of the Shanghai Chenshan
14 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
Fig. 51-58. Mesothorax - 51: Female Phyllium letiranti, dorsal view [SLT coll.]. - 52: Female Phyllium siccifolium, dorsal view [NHMUK]. - 53: Holotype female
Phyllium tobeloense bhaskarai ssp. nov. dorsal view [IMQC]. - 54: Male Phyllium letiranti, dorsal view [SLT coll.]. - 55: Male Phyllium siccifolium, dorsal view
[NHMUK]. - 56: Allotype male Phyllium tobeloense bhaskarai ssp. nov., dorsal view. - 57: Holotype female Phyllium tobeloense bhaskarai ssp. nov. lateral view
of thorax [IMQC]. - 58: Allotype male Phyllium tobeloense bhaskarai ssp. nov. lateral view of thorax [IMQC].
Plant Research Center who identified them as likely a species
of Macaranga (Euphorbiaceae) but without a specimen to
examine a definitive identification is not possible at this time.
Macaranga would not be surprising as this has been recorded
as a phasmid host plant in the past for several other phasmids
(Seow-Choen, 1997). The native people of Morotai call the tree
by the common name “Liwiwi”.
Also accepted as a host plant was Psidium guajava (Myrtaceae)
by individuals collected in the wild and kept in captivity by
Edy Bhaskara (Fig. 101).
Morphological description of the female.
Coloration. — Coloration descriptions are largely based on photographs
of live material sent to the authors from collectors on Morotai. Preserved
specimens with color that was decently retained were also examined to
help describe the spectrum of observed colors.
Two main color forms have been observed, a green form (Fig. 103) and
a yellow/orange form (Fig. 102). In both color forms the antennae are of
a slightly darker color than the rest of the body and with the terminal
three segments with more dense setae and a general darker color than
the other antennae segments. Compound eyes of both forms are
generally of a similar color to that on the head capsule or slightly lighter
in color (in some dried specimens the eyes were a reddish brown but
that is possibly from the drying of the specimen). In the green form, the
serration of the femoral lobes, the margins of the mesothorax, and the
venation of the tegmina are lighter in color than the remaining surfaces,
generally a pale green/yellow or cream color.
In the yellow/orange form, the wing venation is a darker orange than the
tegmina ground color which can be more yellow than orange. In both
color forms the ventral surface of the metacoxae have a clear black
marking, mesocoxae without, cream, yellow, or pale green only.
Morphology
Head. — Capsule \onger than wide (1.1-1.2 times). — Vertex nearly
smooth, with only slight texture and no notable granules except the
prominent posteriomedial tubercle. Frontal convexity stout, only slightly
shorter than the length of the first antennomere, and with a moderate
number of clear setae throughout. — Antennae consisting of nine segments,
with the terminal two segments (VIII and IX) approximately the same
length, and segments IV through VII shorter than the terminal two, and
IV through VII of a uniform length (Fig. 76). — Antennomeres I-VII
sparsely marked with transparent setae (segments I through III with
longer setae than the following segments), and the terminal two
antennomeres are more densely covered in brown setae with fewer
transparent setae. — Compound eyes snug to the head capsule, not
strongly protruding from the head capsule. — Ocelli absent. Antennal
fields wider than the first antennomere but not protruding back farther
than the frontal suture.
Faunitaxys, 7(4), 2019: 1 — 25. 15
Thorax. — Pronotum with anterior and lateral margins that are nearly
straight, lateral margins converging to a straight posterior margin that is
about 60 percent the width of the anterior rim. The pronotum surface
lacks nodes but has prominent furrows giving it a lumpy surface. The
surface has a prominent pit in the center and one along the sagittal plane
near the anterior rim, the remaining furrows are prominent and clearly
visible. The pronotum has a moderate anterior rim, lateral rims are
weak, and the posterior lacks a rim. — Prosternum and the anterior
quarter of the mesosternum with slight granulation. — Metasternum
rather smooth, without prominent granulation. — Mesopraescutum
approximately as long as wide, with lateral rims with five prominent
tubercles rather uniform in size, with occasionally one to two small
nodes between the anterior most tubercles. — Mesopraescutum anterior
rim prominent with a large tubercle in the center with the remainder of
the rim without clear nodes but slightly lumpy, not perfectly smooth
(Fig. 53). — Mesopraescutum crest (excluding the tubercle of the anterior
rim) nearly flat, not prominent, with four small nodes unevenly spaced
along the sagittal plane (Fig. 53). — Mesopraescutum surface lumpy, but
lacking clear nodes. — Mesopleurae straight and evenly diverging; lateral
margin with five to seven major tubercles which are almost always even
in size; occasionally those with five tubercles have one to two smaller
tubercles present (Fig. 53). Face of the mesopleurae smooth, with two
divots, one more prominent divot on the anterior third and one closer to
the center that is faint. — Tegmina slightly variation in length, at the
shortest only reaching the anterior margin of segment VIII and at the
longest reaching just over three quarters the way through VIII.
Alae. — Rudimentary, only about 6mm in length.
Abdomen. — Abdominal segments IT through the anterior two thirds of IV
diverging, posterior third of IV through VII gently converging. — Segments
VII-X converge more acutely than the previous segments, converging
uniformly to a terminal segment that is about as wide as it is long with a broad
rounded apex. — Subgenital plate starts at the anterior margin of segment VIII
and extends three quarters of the way into segment X ending in a fine point,
leaving only the tips of the gonapophyses exposed. — Gonapophyses long,
reaching the posterior margin of segment X (Fig. 65). — Cerci relatively flat
not strongly cupped, with a prominently granular surface and very few short
setae near the apex only.
Legs. — Profemoral exterior lobe smoothly arcing from end to end
without an angle, exterior lobe approximately the same width as the interior
lobe, or slightly thinner and with a granular surface. — Profemoral interior
lobe obtuse angled and marked with five to seven serrate teeth with
generally four to five of these teeth larger than the other smaller teeth and
not perfectly evenly spaced. — Mesofemoral interior lobe smoothly arcs
from end to end and has five to seven serrate teeth distributed on the distal
half. — Interior and exterior lobes are of a similar width, but the exterior lobe
is clearly angled, not smoothly arcing like the interior. — Mesofemoral
exterior lobe with generally only a single small tooth on the distal half of the
arc, rarely an additional smaller tooth is present. — Metafemoral interior lobe
wider than exterior lobe, and arcs end to end and has nine to eleven serrate
teeth pointing distally. — Metafemoral exterior lobe is thin and smooth,
hugging the metafemoral shaft, lacks teeth. — Protibiae lacking an exterior
lobe. — Protibial interior lobe spans the entire length in an evenly
weighted triangle and is about twice the width of the protibiae shaft itself:
— Mesotibiae and metatibiae lacking exterior and interior lobes.
Morphological description of the male.
Coloration. — Most of the body is of a lime green color with little
variation throughout. Ventral of coxae a cream color, without black
markings. The antennae are of a slightly darker green/brown than that
found throughout the body. Protibial interior lobe occasionally with
slight brown markings.
Morphology
Head. — Capsule slightly longer than wide, with a weakly granular
vertex. Frontal convexity stout with a dull point and sparsely marked
with transparent setae. — Antennae consisting of 25 segments
(including the scapus and pedicellus which are marked with few
transparent setae), all other antennal segments with a dense short
covering of darker setae relatively uniform in length throughout all
segments. — Compound eyes large and notably protruding away from
the head capsule. — Ocelli developed clearly between compound eyes.
Thorax. — Pronotum with anterior margin slightly concave and lateral
margins that are straight and gently converging to the posterior margin
that is about 60 percent the width of the anterior rim. Anterior and lateral
margins with moderate rims, posterior margin without a rim and smooth.
Face of the pronotum with moderate furrow on the anterior half and the
surrounding surface without granulation and smooth. — Prosternum with
slight granulation throughout, not highly noticeable. — Mesosternum
surface with more noticeable granules on the anterior. — Metasternum
surface mostly smooth. — Mesopraescutum about as wide as long, with
lateral rims with six to seven tubercles of varying sizes spread nearly
evenly throughout. — Mesopraescutum anterior rim prominent with a
large tubercle in the center with the remainder of the rim smooth (Fig. 56).
— Mesopraescutum crest (excluding the tubercle of the anterior rim) with
three to four small nodes unevenly spaced along the sagittal plane (Fig.
56). — Mesopleura diverging weakly on the anterior but more prominently
on the posterior half; lateral margin with six to seven major tubercles
throughout, with clear gaps between each tubercle with at most two small
nodes in the gaps. — Mesopleural face with two faint divots, one on the
anterior third and one on the posterior third, anterior and posterior of the
face smooth, center of mesopleural face with a wrinkled texture. — Tegmina
reaching half way through abdominal segment I'V or to the posterior margin
on IV.
Alae. — Well developed, reaching segment IX or partly into the anal
abdominal segment.
Abdomen. — Ovular, with segments II through IV diverging, V and
VI parallel or slightly convex, VII through X steadily converging to a
blunt rounded apex. — Anal abdominal segment longer than wide with
lateral margins that converge slowly throughout the majority of the
length and then converge prominently at the terminal third. — Poculum
starts in abdominal segment VIII, with parallel lateral margins, ending
in a broad apex that slightly reaches under segment X. — Cerci are
slightly cupped and exceed the length of segment X, lateral margins
are straight and the surface of the cerci are covered in nodes and short
brown setae throughout. — Vomer broad with nearly straight margins
converging to the apical hook which points upwards into the
paraproct.
Legs. — Profemoral exterior lobe lightly granular and _ slightly
thinner than the profemoral shaft, lobe gently arcing along the entire
profemoral length. — Profemoral interior lobe about as wide as the
shaft and the exterior lobe combined, situated on the distal two thirds
and obtuse, not sharply angled. Profemoral interior lobe with four to
five serrate teeth of similar sizes or with those in the center only
slightly larger. The spacing is uniform except for the middle space
which is slightly greater than the spacing on the sides. — Mesofemoral
exterior lobe about as wide as the shaft, arcing from end to end with
the widest portion weighted slightly off center on the distal end and
with two to four small serrate teeth. — Mesofemoral interior lobe
slightly thinner than the exterior lobe and with less of a bend, and six
to eight serrate teeth on the distal half of the lobe. — Metafemoral
exterior lobe thin and lacking distinct dentition but with a slight rough
texture, hugging femoral shaft. — Metafemoral interior lobe slightly
wider than exterior lobe with ten to twelve serrate teeth throughout the
distal two-thirds of the length. No exterior protibial lobe, interior lobe
extends the entire length in a rounded triangle, with the widest portion
in the center or slightly off center on the distal end and slightly wider
than the shaft width. — Meso- and metatibia simple, lacking lobes.
Description of Egg (Fig. 85 & 86).
The lateral surfaces are flattened and the dorsal and posterior
surfaces are slightly convex, which gives the egg a slightly
rounded ovular appearance. When viewed from the lateral
aspect, the anterior and posterior widths of the capsule are
similar, with the center of the capsule the widest point. When
viewed from the lateral aspect, the anterior, posterior, and
lateral margins are set with numerous rope-like pinnae, each
with a split, slightly hooked apex giving them a barbed
appearance (Fig. 92). Lateral surfaces with textured surface of
circular markings, with 16 to 25 of those circular markings
developed into deeper pits. These pits are in no detectable
pattern and vary from egg to egg. Area around all pits lacking
16 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
Keys to known species of Phyllium from Wallacea
Keys are broken into female, male, and egg morphology due to the marked sexual dimorphism. Female and egg Ph. (Ph.) conlei
and Ph. (Ph.) sp. “Buton” unknown. Egg of Ph. (Ph.) siccifolium unknown. Male of Ph. (Ph.) mamasaense, Ph. (Pu.) keyicum and
Ph. (Pu.) suzukii unknown. Males of Ph. (Ph.) tobeloense tobeloense and Ph. (Ph.) tobeloense bhaskarai ssp. nov. cannot be
morphologically separated and are simply listed together as Ph. (Ph.) tobeloense in the below key.
Female
1. Mesopraescutum distinctly wider than long (>1.7 times) (Fig. 8 & 9)
— Mesopraescutum about as long as wide, or longer than wide (Fig. 51-53)
2. Protibia with an exterior spur near the center (Fig. 7 arrow); profemoral posterior margin distinctly recurved (Fig. 7 line); length
to width ratio of the mesopraescutum approximately | : 3.4 (Fig. 9) Ph. (Pu.) suzukii
— Protibiae lacking exterior lobe; profemoral posterior margin straight (Fig. 6 line); length to width ratio of the mesopraescutum
approximately 1 : 1.8 (Fig. 8) Ph. (Pu.) keyicum
3. Alae developed (Fig. 11)
— Alae rudimentary
4. Mesocoxae or metacoxae with distinct black interior marking (Fig. 46-49); antennae with nine segments
— Mesocoxae and metacoxae without distinct black interior marking (Fig. 50); antennae with ten segments
5. Mesocoxae and metacoxae with distinct black interior marking; exterior lobe of the profemora wide and gently angled with
the greatest width 2.5 to 3 times the width of the profemora shaft (measured at the promixal edge of the interior lobe of the
profemora) (Fig. 30-31)
— Metacoxae with distinct black interior marking, mesocoxae slightly orange or light green, no black interior mark; exterior lobe of
profemora thin and smoothly arcing with the greatest width only 1.5 to 2 times the width of the profemora shaft (measured at the
promixal edge of the interior lobe of the profemora) (Fig. 38-41)
6. Three to five teeth of approximately even size on the interior lobe of the profemora (Fig. 30-31); abdominal segment VIII margin
always almost perfectly straight (Fig. 69 line ) Ph. (Ph.) siccifolium
— Two large teeth and smaller minor teeth between on the interior lobe of the profemora (Fig. 33); abdominal segment VIII margin
always with a smooth bend giving a slight lobe to the segment (Fig. 68 line) Ph. (Ph.) mamasaense
7. Subgenital plate stout, protruding about half the length of sternite X, leaving a significant amount of the gonapophyses exposed
Ph. (Ph.) tobeloense tobeloense
— Subgenital plate long, about three-quarter the length of sternite X, leaving only the tips of the gonapophyses exposed
Ph. (Ph.) tobeloense bhaskarai ssp. nov.
Male
1. Profemoral exterior lobe wider than the interior lobe, strongly angled (Fig. 16)
— Profemoral exterior lobe thinner than the interior lobe, arcing end to end, not strongly angled (Fig. 42-45)
2. Interior lobe of the protibia thin and arcing from end to end without a distinct triangular shape and slender, only about as wide as the
protibial shaft; abdomen long and slender, at its greatest width only about 30% of the total abdomen length (Fig. 23) Ph. (Ph.) conlei
— Interior lobe of the protibia with a distinct triangular shape and wider than the protibial shaft; abdomen broad, with a greatest
width of about 37-50% of the length of the total abdomen
3. Abdomen spade shaped with abdominal segments V through the apex distinctly converging (Fig. 24) Ph. (Ph.) sp. “Buton”
— Abdomen ovoid, with abdominal segments V and VI with parallel margins or only slightly subparallel, only segments VII through
the apex distinctly converging (Fig. 25)
4. Antennae when resting back along the dorsal surface are similar in length or longer than the folded tegmina (Fig. 28)
— Antennae when resting back along the dorsal surface are shorter than the folded tegmina (Fig. 25) Ph. (Ph.) tobeloense
5. Large species 67.0-73.0 mm; antennae apex when resting back along the dorsal surface reaching apex of tegmina or only
exceeding length by one to two segments; mesopraescutum stout (length to maximum width ratio of about 0.8-0.9) and with an
anterior that is distinctly broader than the posterior (Fig. 55) Ph. (Ph.) siccifolium
— Medium species 57.6-62.2 mm; antennae apex when resting back along the dorsal surface exceeding apex of tegmina by four to
five segments; mesopraescutum long and thin (length to maximum width ratio about 1.0-1.2) and with anterior and posterior
margins approximately the same width (Fig. 54) Ph. (Ph.) letiranti
Faunitaxys, 7(4), 2019: 1 — 25. 17
Egg
1. Capsule entirely lacking distinct pinnae
— Capsule with distinct pinnae (most prominent along the operculum rim, if not prominent along the rim, then with short distinct
pinnae along the longitudinal margins of the capsule itself)
2. Lateral margins of the micropylar plate with four to five large pits throughout the length
— Lateral margins of the micropylar plate with eight to nine small pits throughout the length
Ph. (Pu.) suzukii
Ph. (Pu.) keyicum
3. Operculum raised, not flat, lacking long pinnae along the rim (Fig. 81)
— Operculum flat, with long straight pinnae along the rim (Fig. 83-90)
4. Opercular rim with pinnae that are broad and featherlike (Fig. 91), broadest throughout their length
— Opercular rim with pinnae that are thin, ropelike, broadest at the apex (Fig. 92)
5. Lateral surface with short pinnae, notably shorter than the pinnae of the posterior and anterior ends (Fig. 88)
— Lateral surface with rows of long pinnae, of a similar length to the pinnae of the posterior and anterior ends (Fig. 90)
Ph. (Ph.) letiranti
Ph. (Ph.) mamasaense
6. Overall shape somewhat reniform (kidney shaped) (Fig. 83); long pinnae along the margins not fully formed around the entire
capsule, area opposite the microplylar plate and below the operculum bald (Fig. 83)
Ph. (Ph.) tobeloense tobeloense
— Overall shape more symmetrical, ovoid in shape (Fig. 85); long pinnae along the margins fully formed around the entire capsule,
no bald portion opposite of the microplylar plate below the operculum (Fig. 85)
long pinnae, leaving the lateral surfaces without significant
projections. Micropylar plate covering approximately half of
the overall length, situated near the center, not weighted
towards either end of the capsule. Micropylar plate in a thin
teardrop shape, with most of the width thinner than the
micropylar cup. Micropylar cup of moderate size and placed on
the posterior third of the capsule. Operculum nearly circular,
with around ten round impressions encircling the center, and
the outer margin of the operculum has a row of pinnae similar
in length to those found along the other edges of the capsule.
Overall color yellow, tan, or light brown.
Measurements [mm].
— length (including operculum) : 3.8
— length (excluding operculum) : 3.5
— height : 2.1
— width : 2.4
— length of micropylar plate : 2.2
Etymology. — This new subspecies is dedicated to Edy
Bhaskara from Malang, Indonesia who was the first to send us
specimens for this description. May he be thanked for his
numerous collaborations to discover new taxa.
Note on Phyllium female genitalia in relation to
egg morphology
As discussed in depth above, the only easily observed
distinguishing features between the two subspecies of Phyllium
tobeloense are egg morphology and the female subgenital plate
length. This difference in subgenital plate length is likely linked
to the morphological variation observed between the eggs of
the two populations, with subgenital plate length reflecting the
ideal arrangement to hold the egg before it is flung away.
Within the Phylliidae in general, the ratio of subgenital plate
and gonapophyses lengths help to reveal general features of egg
morphology. Members of the bioculatum species group with
eggs characterized by five distinct ribs along the egg (with the
exception of Phyllium giganteum Hausleithner, 1984 which has
3 rounded ribs) have females with shorter subgenital plates and
Ph. (Ph.) tobeloense bhaskarai ssp. nov.
shorter gonapophyses, likely because the eggs can easily be
held with this arrangement or the shorter features aid in the
action of flinging the egg away from the female (Fig. 59). In
contrast, the frondosum species group have short subgenital
plates and longer gonapophyses (Fig. 61 & 62), with eggs
which lack these distinct ribs. In this case the longer
gonapophyses are likely needed to help hold the egg in place
before flinging it. This arrangement of a short subgenital plate
and long gonapophyses is also seen in Phyllium giganteum
(Fig. 60) likely for the same reason, as Ph. giganteum eggs
have stout rounded ribs much like the rounded ribs of the
frondosum species group. Members of the Phyllium (Phyllium)
subgenus generally all have longer gonapophyses and a range
of subgenital plate lengths (presumed to be according to the
ideal length for their specific egg shapes). With eggs of varying
morphologies, the subgenital plate appears to be the feature
most variable among females with differing egg morphologies,
as observed by the two Ph. tobeloense subspecies and their
eggs.
Biogeography and discussion
With nine species of Phyllium currently known from
throughout Wallacea (Fig. 2), several biogeographical trends
can clearly be seen. Particularly the Weber Line which
separates the Papuan subregion species (Ph. suzukii and Ph.
keyicum within the frondosum species group, which is only
found additionally in the Papuan subregion) from the Oriental
elements, particularly illustrative Ph. celebicum, the only
member of the celebicum species group found this far south,
with the others either father north in the Philippines,
Micronesia, or on mainland Asia. The Weber line also
illustrates the likely delineation between Ph. letiranti (west of
the line: Banggai and Sula Islands) from Ph. siccifolium (east
of the line: Buru, Ambon, and Seram Islands).
Of the Lesser Sunda Islands within Wallacea, currently only
Phyllium conlei from Lombok Island is known, with all other
islands devoid of literature and specimen records (Cumming,
Valero & Teemsma, 2018).
To date Phyllium mamasaense has only been recorded from the
type locality of Mamasa, Sulawesi, but with the other
18 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
Fig. 59-69. Fig. 59-67. Genitalia ventral view. - 59: Phyllium abdulfatahi Seow-Choen, 2017 [RC16-039], short subgenital plate and short
gonapophyses. - 60: Phyllium giganteum [RC16-044], short subgenital plate and long gonapophyses. - 61: Phyllium suzukii holotype [SDEI] female short
subgenital plate and long gonapophyses. - 62: Phyllium keyicum [NHMUK 012496951 (Kei Is., 1923)]. - 63: Phyllium siccifolium [FH 1042-1] Seram
Island. - 64: Phyllium tobeloense tobeloense [FH 0657-4] Halmahera Island, Tobelo. - 65: Phyllium tobeloense bhaskarai ssp. nov. holotype female
[IMQC]. - 66: Phyllium tobeloense tobeloense [FH 0657-6] Halmahera Island, Tobelo. - 67: Phyllium tobeloense bhaskarai ssp. nov. allotype male
[IMQC]. Fig. 68-69. Dorsal view of female terminal abdominal segments; lines show the curve of segment VIII, illustrating a consistent feature found
between Ph. mamasaense and Ph. siccifolium. - 68: Holotype Phyllium mamasaense [SDEI]. - 69: Phyllium siccifolium [FH 1042-1].
Sulawesian species Ph. celebicum found throughout the island
it would not be surprising if Ph. mamasaense were also more
widely distributed on the island.
Within Wallacea there are a number of islands currently devoid
of Phylliidae records, some of the largest being from the
eastern Lesser Sunda Islands: Sumbawa, Sumba, Flores, and
Timor. With the island of Timor a rather large island situated
just south of the Phylliidae rich island of Sulawesi, the authors
expected there to be records from Timor as well. However,
after speaking with Chris Reid, Principal Research Scientist at
the Australian Museum in Sydney he is skeptical Phylliidae can
be found on the island. The Australian Museum has been
conducting field work on Timor since 2012 and _ his
observations were as follows, “Most of Timor is dry woodland
on limestone dominated by either Eucalyptus, Casuarina, or
Acacia (now mostly introduced spp), with a bit of sandalwood
etc. Plus a lot of pasture and a lot of weedy arable. The
remaining rainforest is almost entirely dry vine thicket — the
wetter stuff is either converted to coffee or too remote for easy
access or we just didn't get into it. So in general I’d say
Phyllium is unlikely” (pers. comm. Chris Reid).
Undoubtedly, our current knowledge of Wallacean Phylliidae is
still fractional. Of the nine species that have so far become
known from throughout Wallacea, five are only known from a
single sex: Ph. conlei and Ph. sp. ‘Buton’ are both only known
from unique males, and Ph. mamasaense, Ph. keyicum and Ph.
suzukii are only known from females. There is another
questionable celebicum-like species only known from an old
female specimen labelled “Amboina” in the collection of
MNHU, whose identity deserves clarification by additional
Faunitaxys, 7(4), 2019: 1 — 25. 19
Fig. 70-77. Phyllium tobeloense bhaskarai ssp. nov. - 70: Holotype female dorsal [IMQC]. - 71: Paratype female dorsal [FH 1077-1]. - 72: Paratype female
ventral [FH 1077-1]. - 73: Paratype male [FH 1077-22]. - 74: Paratype male [FH 1077-23]. - 75: Dorsal view of terminal abdominal segment (= tergum X) and
cerci, holotype [IMQC]. - 76: Dorsal view of antennae, holotype female [IMQC]. - 77: Dorsal view of antennae, allotype male [IMQC].
CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
s3) ~
— ~
-
Fig. 78-80. Macaranga sp. (Euphorbiaceae) with wild Phyllium (Phyllium) tobeloense bhaskarai ssp. nov. on Morotai Island (© Alim Yasin).
- 78: Macaranga sp., host plant. - 79: Subadult female on Macaranga sp. - 80: Adult male on Macaranga sp.
material (see discussion on PA. celebicum). Also, the authors
are aware of at least one further still undescribed Phyllium
species which is only known from a photograph. It is very
much hoped that the present paper will encourage collectors to
look out for Phylliidae throughout Wallacea to obtain fresh
specimens that can help broaden our knowledge of these
fascinating insects from this particular region so well known
for its high biodiversity and high degree of endemism.
Acknowledgments
The authors thank René Limoges, entomological technician at
the Montreal Insectartum for taking many photos for this work,
as well as for many professional courtesies. Thank to Vadim
Voitsekhovskii (Bali, Indonesia) for sending a series of
specimens for this study. A special thanks to Alim Yasin
(Morotai, Indonesia) who sent to us an extra 50 males and 50
Faunitaxys, 7(4), 2019: 1 — 25. 21
45
—_—
one ,
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ae
4
—
>
a ee te | rs
ie:
ARs
; - t
ee ee
— She
Weta f
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Biicp US
o1 Si) Ra A
.
&
Fig. 81-92. Phyllium eggs. Fig. 81, 83, 85, 87, 89 lateral view, m
87
a eS —— cu
Sa At. Ss Se
WF. > s kes ... a
Q Pr ed — a
> ae
‘ ae aw" on
4 my &
92
micropylar plate. - 81 & 82: Phyllium celebicum. - 83 & 84: Phyllium tobeloense tobeloense. - 85 & 86: Phyllium tobeloense bhaskarai ssp.
nov. paratype. - 87 & 88: Phyllium letiranti paratype. - 89 & 90: Phyllium mamasaense (© Gré8er D.). Fig. 91-92. Pinnae from the opercular
rim. - 91: Featherlike pinnae, frills throughout length, wide throughout length, apex not widest portion, Phyllium letiranti. - 92: Ropelike pinnae,
stalk without frills, broadest portion the apex, Phyllium tobeloense tobeloense. Scale bar = 2 mm.
females that allowed us to deposit many paratypes in various
museums and who also sent us photos of wild specimens and
the habitat. Thanks to Maxime Ortiz (France) who sent us eggs
that have been used in this work. Thanks also to Binjie Ge
(Shanghai Chenshan Plant Research Center and Gilles Vincent,
Special advisor of the Executive President of the Chenshan
Botanical Garden-Shanghai, China) for the identification of the
host plant. Thank you to Jim Berrian (SDNHM, San Diego) for
years of support and access to the museum’s camera. Thank
you to Tim Van molle (Rupelmonde, Belgium) for allowing us
to use photos of his beautiful Phyllium tobeloense color forms
and Detlef GroBer (Berlin, Germany) for allowing us to use his
photos of Phyllium mamasaense eggs in this work. Sigetake
Suzuki (Hokkaido, Japan) kindly provided various specimens
from Sulawesi and helpful information. We thank Edy
Bhaskara (Malang, Java, Indonesia) for providing the
specimens of Phyllium tobeloense bhaskarai ssp. nov. and Ph.
tobeloense tobeloense for description as well as pictures of live
specimens of the new subspecies. Thanks are also due to the
curators of various museums for access to the corresponding
collections that contain material listed in the present paper and/
or providing photographs of specimens: Dr. Michael Ohl and
Ulrike Dorandt (MNHU, Berlin), Dr. Susanne Randolf and
Prof. Ulrike Aspé6ck (NHMW, Vienna), Dr. Caroline Pepermans
(RMNH, Leiden) and Hans Mejlon (UUZM, Uppsala). Mandy
Schréter and Stephan Blanke (SDEI, Senckenberg German
Entomological Institute Miincheberg) for photographs of
specimens used in this work. Judith Marshall and Benjamin
Price (NHMUK, London) for processing the loan of the
Phyllium keyicum specimen despite the lack of an Orthopteroid
collection curator. To conclude we of course thank our two peer
reviewers who gave us valuable feedback and suggestions on
how to improve this text, thank you Philip E. Bragg
(Nottinghamshire, United Kingdom) and Dr Allan Taylor
(Quebec, Canada).
CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
Fig. 93-95. Live Phyllium celebicum captive reared F1-generation. - 93: Male/female pair. - 94: Male. - 95: Female.
Faunitaxys, 7(4), 2019: 1-25.
Fig. 96-98. Live Phyllium letiranti captive reared (© Maxime Ortiz). - 96: Yellow form female. - 97: Yellow form female, detail of head, thorax, and
profemoral lobes. - 98: Phyllium letiranti Males.
24 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
Fig. 99-100. Live Phyllium tobeloense tobeloense captive reared female (© Tim Van molle). - 99: Red color form. - 100: Yellow and green color forms.
Fig. 101-103. Live Phyllium tobeloense bhaskarai ssp. nov. - 101: Captive adult pair feeding on Psidium guajava (Myrtaceae) (© Alim Yasin). - 102: Orange
form female, paratype (© Alim Yasin). - 103: Green form female, paratype (© Edy Bhaskara).
Faunitaxys, 7(4), 2019: 1 — 25. 25
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Cumming R. T., Le Tirant S. & Hennemann F. H., 2019. — Révision des Phyllium Illiger, 1798 de Wallacea (Indonésie), avec la description
d’une nouvelle sous-espéce de l’ile Morotai (Phasmatodea: Phylliidae: Phylliinae). Faunitaxys, 7(4) : 1-25.
Les Phylliidae de Wallacea sont révisés. La répartition des espéces est précisée. Une nouvelle sous-espéce de Phyllium
(Phyllium) tobeloense Gréker, 2007 est décrite de l’ile de Morotai, Phyllium (Phyllium) tobeloense bhaskarai Cumming, Le
Tirant et Hennemann ssp. nov. dont l’identification est basée principalement sur la morphologie distincte des ceufs. Enfin une
clé de tous les Phyllium connus de Wallacea est présentée permettant une identification rapide.
Mots-clés. — Phasmatodea, Phasmida, Phylliinae, Phylliini, Phyllium, Wallacea, Indonesia, Morotai, tobeloense, bhaskarai, description,
espéce nouvelle, distribution, biogeographie.
Faunitaxys
Volume 7, Numero 4, Mars 2019
SOMMAIRE
Révision du genre Phyllium Illiger, 1798 de Wallacea, avec la description d’une nouvelle sous-
espéce de l’ile Morotai (Phasmatodea: Phylliidae: Phylliinae).
Royce T. Cumming, Stéphane Le Tirant & Frank H. Hennemann
CONTENTS
Review of the Phyllium Illiger, 1798 of Wallacea, with description of a new subspecies from
Morotai Island (Phasmatodea: Phylliidae: Phylliinae).
Royce T. Cumming, Stéphane Le Tirant & Frank H. Hennemann
Mlustration de la couverture: Forét tropicale de Morotai.
Crédit:
Fig. 2 © (Google Earth : Image Landsat).
Fig. 3-10-11-12-13-14-15-16- 31-34-36-38-4 1-43-45-47-48-63-64-66-69-7 | -74-8 1-84-89-9 1-93-95 © Hennemann F.H., Germany.
Fig.-29-32- 49-52 © Natural History Museum, United Kingdom.
Fig. 5-7-9-17-21-22-24-25-26-28-33- 46-61-68 © Senckenberg Deutsches Entomologisches Institut, Miincheberg, Germany.
Fig. 4-6-8-18-27-37-42-5 1-53-54-56-57-58 -62-65-67-70-75-76-77-85-86-88 © Limoges R., Insectarium de Montréal, Québec, Canada.
Fig. 20-30 © Uppsala University Zoological Museum, Linnean collection, Uppsala, Sweden.
Fig. 23-44 © Zoologische Staatssammlung, Munich, Germany.
Fig. 50-96-98 © Ortiz M., France.
Fig. 59-60 © Cumming R.T., U.S.A.
Fig. 78-80-101-102 & couverture © Yasin A., Indonesia.
Fig. 89-90 © GréBer D., Germany.
Fig. 99-100 © Van molle, T, Belgium.
Fig. 103 © Bhaskara E., Indonesia.
Publié par |’ Association Frangaise de Cartographie de la Faune et de la Flore (AFCFF)
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